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The Case for the Entourage Effect and Conventional Breeding of Clinical Cannabis: No “Strain,” No Gain

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The Case for the Entourage Effect and Conventional Breeding of Clinical Cannabis: No “Strain,” No Gain fpls-09-01969 January 8, 2019 Time: 15:54 # 1 PERSPECTIVE published: 09 January 2019 doi: 10.3389/fpls.2018.01969 The Case for the Entourage Effect and Conventional Breeding of Clinical Cannabis: No “Strain,” No Gain Ethan B. Russo* International Cannabis and Cannabinoids Institute, Prague, Czechia The topic of Cannabis curries controversy in every sphere of influence, whether politics, pharmacology, applied therapeutics or even botanical taxonomy. Debate as to the speciation of Cannabis, or a lack thereof, has swirled for more than 250 years. Because all Cannabis types are eminently capable of cross-breeding to produce fertile progeny, it is unlikely that any clear winner will emerge between the “lumpers” vs. “splitters” in this taxonomical debate. This is compounded by the profusion of Cannabis varieties available through the black market and even the developing legal market. While labeled “strains” in common parlance, this term is acceptable with respect to bacteria and Edited by: viruses, but not among Plantae. Given that such factors as plant height and leaflet Giuseppe Mandolino, Council for Agricultural and width do not distinguish one Cannabis plant from another and similar difficulties in Economics Research, Bologna, Italy defining terms in Cannabis, the only reasonable solution is to characterize them by their Reviewed by: biochemical/pharmacological characteristics. Thus, it is best to refer to Cannabis types Gianpaolo Grassi, as chemical varieties, or “chemovars.” The current wave of excitement in Cannabis CREA-CIN Rovigo, Italy Raffaella Pergamo, commerce has translated into a flurry of research on alternative sources, particularly Council for Agricultural and yeasts, and complex systems for laboratory production have emerged, but these Economics Research, Rome, Italy presuppose that single compounds are a desirable goal. Rather, the case for Cannabis *Correspondence: Ethan B. Russo synergy via the “entourage effect” is currently sufficiently strong as to suggest that one [email protected]; molecule is unlikely to match the therapeutic and even industrial potential of Cannabis [email protected] itself as a phytochemical factory. The astounding plasticity of the Cannabis genome Specialty section: additionally obviates the need for genetic modification techniques. This article was submitted to Keywords: cannabis, cannabinoid, marijuana, hemp, genomics, genetically modified organism, Plant Breeding, tetrahydrocannabinol, cannabidiol a section of the journal Frontiers in Plant Science Received: 31 October 2018 INTRODUCTION: DEFINING TERMS Accepted: 19 December 2018 Published: 09 January 2019 Earlier data on taxonomy of Cannabis was previously reviewed (Russo, 2007), which will be Citation: herein summarized and supplemented. Cannabis is a dioecious annual of the Cannabaceae Russo EB (2019) The Case family which traditionally includes hops, Humulus spp. Alternatively, Cannabis has also been for the Entourage Effect and Conventional Breeding of Clinical assigned to Moraceae, Urticaceae, or even in the Celtidaceae families on the basis of chloroplast Cannabis: No “Strain,” No Gain. restriction site maps (Weigreffe et al., 1998), and chloroplast mat K gene sequences (Song Front. Plant Sci. 9:1969. et al., 2001). More recently, the Cannabaceae have subsumed eight genera: Celetis, Pteroceltis, doi: 10.3389/fpls.2018.01969 Aphananthe, Chaetachme, Gironniera, Lozanella, Trema, and Parasponia, comprising 170 odd Frontiers in Plant Science| www.frontiersin.org 1 January 2019| Volume 9| Article 1969 fpls-09-01969 January 8, 2019 Time: 15:54 # 2 Russo Conventional Breeding of Cannabis species (McPartland, 2018), a finding supported by genetic The Cannabis species controversy, Cannabis sativa vs. indica analysis of four plastid loci (Yang et al., 2013). Current vs. afghanica, has continued unabated to the current day research on fossil pollen samples associated with the ecological with impassioned arguments advanced by the protagonists associations of Cannabis with steppe companion species (Clarke and Merlin, 2013, 2016; Small, 2015; McPartland (Poaceae, Artemisia, Chenopodiaceae), and Humulus (hops) with and Guy, 2017; Small, 2017). This author, having been forest genera (Alnus, Salix, Populus), have established that on every side of the issue at one time or another, has although Cannabis seems to have originated in the Tibetan chosen to eschew the irreconcilable taxonomic debate as Plateau at least 19.6 million years ago, it has also been indigenous an unnecessary distraction (Piomelli and Russo, 2016), and to Europe for at least a million years (McPartland et al., 2018), rather emphasize that only biochemical and pharmacological and refuted the conventional wisdom that this “camp follower” distinctions between Cannabis accessions are relevant. In was brought there by man. his recent seminal review, McPartland agreed, “Categorizing The species assignation of Cannabis itself is fraught with Cannabis as either ‘Sativa’ and ‘Indica’ has become an exercise in great debate. Cannabis sativa, meaning “cultivated Cannabis,” futility. Ubiquitous interbreeding and hybridization renders their was so named by Fuchs, among others, in 1542 (Fuchs, distinction meaningless.” (McPartland, 2018) (p. 210). 1999), an assignation 211 years before the systematization of An additional non-sensical nomenclature controversy botanical binomials Linnaeus in his Species Plantarum (Linnaeus, pertains in common parlance to Cannabis “strains,” an 1753). Lamarck subsequently suggested Cannabis indica, a more appellation that is appropriate to bacteria and viruses, but diminutive intoxicating Indian plant from India, as a separate not plants (Bailey and Bailey, 1976; Usher, 1996; Brickell et al., species (Lamarck, 1783). The issue has remained unresolved 2009), especially so with Cannabis where the chemical variety, in the subsequent centuries with two opposing philosophies. abbreviated “chemovar” is the most appropriate appellation Ernest Small has championed the single species concept (Small (Lewis et al., 2018). and Cronquist, 1976). Polytypic treatments of Cannabis also gained adherents (Schultes et al., 1974; Anderson, 1980) on morphological criteria suggesting separation of Cannabis THE CANNABIS GENOME AND sativa L. Cannabis indica Lam. and Cannabis ruderalis Jan., ALTERNATIVE HOST BIOCHEMICAL a scheme supported by systematic chemotaxonomy. Principal PRODUCTION component analysis (PCA) of 157 Cannabis accessions from around the world assessed allozyme frequencies at 17 gene 2011 was a landmark year for Cannabis genomics, as Medical loci suggested a split (Hillig, 2005b). “Sativa” gene pools from Genomics and Nimbus Informatics issued an online report on eastern European ruderal samples were linked to narrow- the complete 400 million base-pair genomic sequence, which was leaflet European and Central Asian fiber and seed plants, shortly joined by a draft genome and transcriptome (van Bakel while an “indica” grouping encompassed Far Eastern seed et al., 2011). and fiber plants and drug plants with broad-leaflets from This development sparked prominent publicity and most of the rest of the world, along with wild accessions controversy as to what it might portend. Whereas, the human from the Indian subcontinent. Central Asian roadside samples genome was analyzed some 20 years earlier, the implications for (Cannabis ruderalis) were thought to represent a third group. Cannabis were subject to great speculation. Gas chromatography (GC) and starch-gel electrophoresis studies The news catalyzed a flurry of new research, but considerable also suggested species separation of sativa and indica (Hillig and progress had already been achieved in applied Cannabis genetics. Mahlberg, 2004). The identification and synthesis of 19-tetrahydrocannabinol Agronomic factors in 69 samples suggested inclusion of (THC) was accomplished in Israel 1964 (Gaoni and Mechoulam, eastern hemp and drug plants in Cannabis indica (Hillig, 1964), but it was not until much later before successful cloning 2005a), a division supported by fragment length polymorphisms of its biosynthetic enzyme, tetrahydrocannabinolic acid synthase (Datwyler and Weiblen, 2006). (THCA synthase) (Sirikantaramas et al., 2004; Figure 1). Enzyme More recently, PCA seemed to point to terpenoid content crystallization followed (Shoyama et al., 2005). Cannabidiolic as the most convincing distinguishing chemotaxonomic markers acid synthase, which catalyzes cannabidiolic acid (CBDA), the between putative sativa and indica species (Elzinga et al., 2015). precursor of cannabidiol (CBD), had been previously identified Similarly, PCA was felt to separate drug Cannabis from hemp and produced in pure form (Taura et al., 1996; Figure 1). (Sawler et al., 2015). A recent study demonstrated demarcation These developments stimulated additional findings, including the of Cannabis drug from hemp accessions via genotyping of archeological phytochemical discovery of THCA synthase in a 13 microsatellite loci across the genome, not merely genes 2700 year old Cannabis cache from a tomb in Central Asia along affecting cannabinoid or fiber production (Dufresnes et al., with two previously unreported single nucleotide polymorphisms 2017). Professor Giovanni Appendino has reported the presence (SNPs) in the enzyme’s gene sequence (Russo et al., 2008). of the cis-19-THC stereo-isomer only in the hemp accessions By 2011, the enzymes for the production of the major (Giovanni Appendino, personal communication). However, phytocannabinoids had been identified.
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